Table Of ContentThe symposium on Tropical Forests: Botanical dynamics, speciation, and
diversity was held at Aarhus University on the occasion of the twenty-fifth
Anniversary of the Botanical Institute. The Danish Natural Science
Research Council, the Research Foundation of Aarhus University, Aarhus
Olie, and DANIDA, are hereby acknowleged for making the event
possible.
Tropical Forests
Botanical Dynamics, Speciation and Diversity
Edited by
L. B. Holm-Nielsen, I. C. Nielsen and H. Balslev
Botanical Institute
Aarhus University
ACADEMIC PRESS
Harcourt Brace Jovanovich, Publishers
London • San Diego • New York
Boston • Sydney • Tokyo • Toronto
ACADEMIC PRESS LIMITED
24-28 Oval Road
London NW1 7DX
United States Edition published by
ACADEMIC PRESS INC
San Diego, CA 92101
Copyright © 1989 by
ACADEMIC PRESS LIMITED
Second Printing 1990
All Rights Reserved
No part of this book may be reproduced in any form by photostat,
microfilm, or any other means, without written permission
from the publishers
British Library Cataloguing in Publication Data
is available
This book is printed on acid-free paper
ISBN 0-12-353550-6
Printed in Great Britain by T. J. Press (Padstow) Ltd.,
Padstow, Cornwall
Foreword
Last year, I was greatly disappointed to be unable to attend the
Tropical Forest Symposium at Aarhus University. Now that I have the
papers in front of me, I am even more sorry that the Symposium coincided
with the time of my international move! This is a wonderful, up-to-date
assessment of dynamics, speciation and diversity in tropical forests. It is
also a worthy tribute to the long and distinguished career of Professor
Larsen, whose vision built the Botanical Institute that celebrated its
twenty-fifth anniversary with this Symposium. Already, in the
comparatively short life of the Botanical Institute of Aarhus University, its
students have become leaders in the international botanical community
and have produced a large number of useful and significant publications.
Tropical forests are disappearing at an alarming rate. To provide a
rationale for their conservation and sustainable utilisation, we must
understand their complexity, interactions and dynamics. This is not just
another volume that proclaims the destruction of forests; it provides the
data on which to base conservation and utilisation programmes.
The section on forest dynamics discusses both the physical features
such as soil, landscape and water, and the patterns of tree distribution in
the forest. These chapters show that, over the last 25 years, we have really
begun to understand forest structure and composition because we have data
from so many fields such as tree architecture, gap-phase dynamics,
canopy structure, etc. The forest is now known to be much more dynamic
than was thought by earlier workers who emphasized long-term stability.
The sections on diversity and speciation are largely written by
systematists. It is most encouraging to see the many ecological and
theoretical aspect that are emerging from their work. The combination of
an ecological and taxonomic approach has led to many useful ideas about
species diversity and speciation. This volume is a fine example of the way
in which existing, but still far from complete, data are being synthesised to
enable us to understand, not just the defination of each of the many tropical
forest species, but also to understand how this remarkable species diversity
is put together into the dynamic and evolving ecosystem that we know as
tropical rain forest. It presents a strong case for the health and value of
contemporary tropical systematics. We just need a larger workforce.
This underlines the increased importance of the few remaining institutes
that are training tropical systematists. This volume will continue to keep
the Botanical Institute of Aarhus University in the forefront of tropical
forest studies.
Ghillean T. Prance, Royal Botanic Gardens, Kew.
ν
Organizing Committee
L. B. Holm-Nielsen, I. C. Nielsen, and H. Balslev
Symposium on
Tropical Forests: Botanical dynamics, speciation and
diversity.
held at Aarhus University, 8-10 August 1988
Session 1. Dynamics - Chairmen P. S. Ashton and P. Windisch
Session 2. Speciation - Chairmen B. ter Welle and Hu Chi-ming
Session 3. Diversity - Chairmen 0. Hamann and B. Nordenstam
vi
Contributors
Andersson, L. Institut for Systematisk Botanik, University of Goteborg,
Carl Skottsbergsgata 22, S-413 19 Goteborg, Sweden (p. 173)
Ashton, P. S. The Arnold Arboretum of Harvard University, 22 Divinity
Avenue, Cambridge, MA 02 138, USA (p. 239)
Balslev, H. Botanisk Institut, Aarhus University, Nordlandsvej 68, DK-
8240 Risskov, Denmark (p. 287)
Barthelemy, D. Laboratoire de Botanique, Institut Botanique, 163 rue A.
Broussonet, F-34000 Montpellier, France (p.89)
Bruenig, E. F. Institut fur Weltforstwirtschaft, University of Hamburg,
Leuschnerstrasse 91, D-2050 Hamburg 80, Federal Republic of
Germany (p. 75)
Castroviejo, S. Real Jardin Botanico, Plaza de Murillo 2, 28014 Madrid,
Spain (p. 347)
Chen Z-y. South China Institute of Botany, Academia Sinica, Guangzhou,
Wushan, Peoples Republic of China (p. 185)
Dransfield, J. Herbarium, Royal Botanic Gardens, Kew, Richmond,
Surrey, TW9 3AB England, UK (p. 153)
Edelin, C. Laboratoire de Botanique, Institut Botanique, 163 rue A.
Broussonet, F-34000, Montpellier, France (p. 89)
Feuillet, C. Centre ORSTOM de Cayenne, B. P. 165, 97323 Cayenne Codex,
French Guiana (p. 311)
Geesink, R. Rijksherbarium, Rapenburg 70-74, 2311 EZ Leiden, The
Netherlands (p. 135)
Gentry, Α· H. The Missouri Botanical Garden, P.O. Box 299, St. Louis,
MO 63166-0299, USA (p. 113 )
Halle, F. Laboratoire de Botanique, Institut Botanique, 163 rue A.
Broussonet, F-34000 Montpellier, France (p. 89)
Hansen, B. Botanisk Museum, University of Copenhagen, Gothersgade
130, DK-1123 Copenhagen K, Denmark (p. 201)
Hartshorn, G. S. World Wildlife Fund, 1250 24th Street, NW,
Washington DC 20037, USA (p. 65)
Haynes, R. R. Department of Biology, University of Alabama,
Tuscaloosa, AL 35486, USA (p. 211)
Holm-Nielsen, L. Β· Botanisk Institut, Aarhus University, Nordlandsvej
68, DK-8240 Risskov, Denmark (p. 211)
Huang, Y-w. Institut fur Weltforstwirtschaft, University of Hamburg,
Leuschnerstrasse 91, D-2050 Hamburg 80, Federal Republic of
Germany (p. 75)
Huber, O. C. V. G. and Instituto Venezolano de Investigaciones Cienti-
ficas (I. V. I. C), Apartado 80405, Caracas 1080-A, Venezuela (p. 271)
vii
Irion, G. Forschungsinstitut Senckenberg, Abteilung fur Meeresgeologie
und Meeresbiologie, Schleussenstrasse 39a, D-2940 Wilhelms-
haven, Federal Republic of Germany (p. 23)
Iwatsuki, K. Botanical Gardens, University of Tokyo, Hakusan, Tokyo
112, Japan (p. 193)
Junk, W. J, Max Planck-Institut fur Limnologie, Arbeitsgruppe fur
tropischer Okologie, P. 0. Box 165, D-2320 Plon, Federal Republic of
Germany (p. 47)
Kornet, D. J. Instituut voor Teoretische Biologie, University of Leiden,
Groenhovenstraat 5, 2311 BT Leiden, The Netherlands (p. 135)
Larsen, K. Botanisk Institut, Aarhus University, Nordlandsvej 68, DK-
8240 Risskov, Denmark (p. 339)
Luteyn, J. L. The New York Botanical Garden, Bronx, NY 10458-5126,
USA (p. 297)
Mori, S. A. The New York Botanical Garden, Bronx, NY 10458-5126, USA
(p. 319)
Nielsen, I. C. Botanisk Institut, Aarhus University, Nordlandsvej 68,
DK-8240 Risskov, Denmark (p. 355)
Oldeman, R. A. A. Institute of Silviculture and Forest Ecology,
Agricultural University, P.O. Box 342, 6700 AN, Wageningen, The
Netherlands (p. 3)
Ortiz-Crespo, F. US-AID, American Embassy, Quito, Ecuador (p. 335)
Polhill, R. M. Herbarium, Royal Botanic Gardens, Kew, Richmond,
Surrey, TW9 3AB, England, UK (p. 221)
Ramella, L. Conservatoire et Jardin Botaniques, Case postale 60, CH-1192
Chambesy/GE/Switzerland (p. 259 )
Rasanen, M. Department of Biology, University of Turku, SF-20500
Turku, Finland (p. 35)
Raven, P. H. The Missouri Botanical Garden, P.O. Box 299, St. Louis, MO
63166, USA (p. 365)
Renner, S. S. Botanisk Institut, Aarhus University, Nordlandsvej 68,
DK-8240 Risskov, Denmark (p. 287)
Salo, J. Department of Biology, University of Turku, SF-20500 Turku,
Finland (p. 35)
Shukla, V· K. S. Analytical Research and Development, Aarhus
Oliefabrik A/S, P.O. Box 50, DK-8100 Aarhus C, Denmark (p. 355)
Spichiger, R. Conservatoire et Jardin Botaniques, Case postale 60, CH-
1292 Chambesy/GE/Switzerland (p. 259)
Sumithraarachchi, D. B. Royal Botanic Gardens, Peradeniya, Kandy, Sri
Lanka (p.253)
Swaine, M. D. Department of Botany, University of Aberdeen, St. Machar
Drive, Aberdeen AB9 2UD, Scotland, UK (p. 101)
viii
Preface
The botany of tropical forest has been a focal point of the research and
teaching of the Botanical Institute since its establishment at Aarhus
University 25 years ago. We have often felt the need for more knowledge
about the diverse tropical ecosystems, for example when we saw the same
meager data used to redundency in the argument for the conservation of
some of these areas. The temperate areas of the world have not been
developed in a sustainable way but we may be learning the lesson. The
tropical forest still houses an unmatched biological richness. In order to
prevent the total destruction of these areas the scientific community must
be urged to produce and communicate further knowledge about them.
As Raven points out in the concluding remarks of this volume 150
000 species of higher plants in the tropics are being studied by only a few
hundred botanists. These species and their potential uses are not likely to
be studied before many of them become extinct. Training of taxonomists
has been neglected in many countries, perhaps because many schools of
taxonomists have isolated themselves from the adjacent disciplines. We
have tried to avoid this in Aarhus and therefore we wanted to expand the
theme of the symposium on Tropical Forest held in Aarhus in August 1988
to include dynamics and diversity aspects. It is our opinion that some of
the major tasks for biologists in years to come will be to achieve better
understanding of the importance of such concepts as dynamics and
diversity of ecosystems. It must be stressed, however, that without further
basic studies of the speciation and the taxonomy of the organisms of the
systems such new understanding is not likely to be achieved. The
research at the Botanical Institute has been directed both towards the
taxonomy of tropical plants, the floristics of selected areas and, more
recently, towards the tropical ecosystems. We have furthermore developed
applied research in some related areas (cf. Shukla and Nielsen, this
volume) and have been engaged in development projects such as
reforestation of the Andean highlands with local species.
The Symposium was arranged to bring together a comprehensive but
not too large group of people from forestry institutes, ecological institutes,
private industry, botanical gardens, museums and universities on the
occasion of the twenty-fifth anniversary of the Botanical Institute. The 144
participants represented 24 countries. Papers were presented orally and as
posters. The 56 abstracts are published in Skov and Barfod, (1988), AAU-
Reports 18, 1-46. The present volume includes the written version of the
oral presentations.
We wish to acknowledge the help of the session chairmen P. S.
ix
Ashton, Ο. Hamann, Hu Chi-ming, B. Nordenstam, B. ter Welle, and
Paulo Windisch who made the program move smoothly, and to F. Ortiz-
Crespo who gave a stimulating morning appeal (Ortiz, this volume) about
the situation of the tropical forests as seen from the point of view of the
people living in the tropical countries.
The papers were rewied by a number of colleauges with special
knowledge of the topics treated. Their promt and critical comments were
most helpful to our editing, and we are most grateful to them: L.
Andersson, P. S. Ashton, A. Barfod, B. Boom, V. A. Funk, A. H. Gentry,
R. Haase, O. Hamann, R. R. Haynes, N. Jacobsen, J. A. Korstgard, J.
Kress, J. Kuijt, K. Larsen, J. L. Luteyn, Τ. V. Madsen, U. Molau, S. A.
Mori, R. A. A. Oldeman, K. Rahn, T. Ray, S. S. Renner, M. Richardson,
K. Sand-Jensen, W. G. Sombroek, K. Torssell, D. Wasshausen, C.
Westerkamp, and B. 011gaard. Special thanks go to S. Churchill who
read through all the papers.
The scientific and technical editing of this book was carried out at
the Botanical Institute, Aarhus University. Without the outstanding help of
the staff the completion of this task within few months would not have been
possible. The editors appreciate the fine artwork by A. Sloth and K. Tind,
and the word processesing and layout by A. Thygesen and B. H0jstr0m.
F. N0rgaard assisted in the computing and A. Boyd was linguistic
consultant.
We wish to emphasize that although we have placed the individual
papers in a context, the full story about the dynamics, speciation and
diversity of the tropical forests is not told. We have found that each of the
published papers in its own way provides us with further ideas about the
tropical forests. We have intended to bring people and thoughts together
which in the further work with these important and exciting forests will
fertilize the creation and transmission of new knowledge to the benefit of
science and the tropical forests.
Lauritz B. Holm-Nielsen, Ivan Nielsen and Henrik Balslev
χ
Introduction
It used to be agreeable to many scientists that at least some areas of tropical
forests existed under relatively stable climatic conditions during the
geologic history in which the Angiosperms evolved.
This myth of the forest ecosystems being stable climaxes has now
been challenged. The present knowledge about the dynamics of these
systems and the evolution of the landscapes in which they occur now
justifies the richness in niches and diversity. The understanding of the
tropical forests as high-diversity systems help us explain the apparent
stability often observed. Further studies will without doubt provide science
with new understanding of the theory and philosophy of diversity as a
concept, and of dynamics as a stabilizing element in nature
In his introductory paper to the section on dynamics in tropical
forests Oldeman presents the hypothesis of ecological interference and the
linkage to the architecture of forest mosaic, the building and the dynamics
of forest eco-units, and the distribution patterns of habitats. The geology
and geomorphology of some regions are about to be better understood (Irion
and Salo), and theories on the history of the Amazon Basin, such as the
refugium theory, presented by Haifer in 1974 must be revised accordingly.
The ecological dynamics caused by rythmic flooding of extended areas in
the Amazon Basin (Junk), is an example of natural dynamic conditions
which may have influenced diversification of the forests and the
speciation processes during the Pleistocene. The small scale dynamics, of
natural and man-made gap structures (Hartshorn) and of canopy
structure as seen during the regeneration or succession resulting from
sporadic events such as forest burning caused by lightning strikes
(Bruenig and Huang) are important features for the understanding of the
forests. The bioarchitectural concept employed by Hall6 and collaborators
(Barthetemy et al.) may prove to be a useful tool in the further and detailed
studies of the structure of tropical forests. The concluding paper by Swaine
draws the attention to some of the research which is needed in order to get
closer to an understanding of the dynamics of tropical forests.
The second chapter on the speciation of tropical plant groups is
introduced by Gentry's review. Though his theory of explosive evolution,
examplified from Cerro Centinela in western Ecuador, where he suggests
that species evolved during 15 years of isolation, may be considered only a
challenging theory, it provides us with new ideas, and reminds us that our
own prejudices may be the major obstacle in the search for further
understanding. The theory of entropy explained and examplified from
Malesian legumes (Geesink and Kornet) provides a philosophical
xiii
